Phosphorus mobility in interstitial waters of sediments in Lake Kinneret,Israel

Monthly samples of sediment cores from maximum depth ( 42 m) in Lake Kinneret were taken from May 1988 until January 1989. The chemical composition of the interstitial and overlying water was investigated with respect to phosphate, Fe²⁺, Fe³⁺, Ca²⁺, alkalinity and electric conductivity. pH, pH₂S and pe (electron-activity) were measured by microelectrodes inserted directly into the sediment core immediately after sampling.Ion activity products of vivianite, siderite, ironsulfides, Ca-P complexes and Ca-P solid phases were calculated; in addition, Ca/P ratios for the overlying and pore water were obtained by using the potential diagram technique. Despite the fact that anoxic conditions prevail for most of the year, no control of phosphate solubility by a Fe-P relationship could be found. Determination of IAPs, together with calculated molar Ca/P-ratios, suggests that hydroxyapatite as well as surface complexes like dicalciumphosphate are the solubility-controlling species in pore water. For the overlying water a Ca₃(HCO₃)₃PO₄ surface complex is assumed to fix the phosphorus, accompanied by a subsequent transformation of the bound P into apatite.     

Sulphate reduction in the hypolimnion and sediments of Lake Kinneret, Israel

1. Lake Kinneret is a warm (13–30°C) monomictic lake. Between January and June a heavy annual bloom of the dinoflagellate Peridinium gatunense dominates phytoplankton biomass (250 g m^?2). At the beginning of the summer, degradation and decomposition of the Peridinium biomass occurs, serving as a trigger for intense sulphate reduction in the hypolimnion and sediments. 2. The rates of sulphate reduction in the sediments varied seasonally from 12 to 1700 nmol SO₄.^?2 reduced cm^?3 day^?1 in December and July, respectively. The availability of organic matter and sulphate is high in June after the crash of the Peridinium bloom and the beginning of stratification and is lowest in December before overturn. 3. Sulphate concentrations in the hypolimnion range between 0.52 mM and 0.20 mM during mixing (January-April) and before overturn (December), respectively. The depletion in sulphate in the hypolimnion is stoichiometrically correlated to the increase in sulphide. The lake is not depleted of sulphate at any time, so the sulphate reduction process in Lake Kinneret is not limited by sulphate concentrations except in the sediments just before overturn.     

Metalimnic Layer in Lake Kinneret,Israel

Summary Chemical and biological investigations of Lake Kinneret showed that reducing conditions prevailling in summer in the metalimnion cause an early and intense denitrification process accompanied by the release of dissolved Fe and Mn. These conditions are favourable to the development of sulfo-bacteria and blue green algae. The role of this layer is discussed.     

Sulfate reduction processes in sediments at different sites in Lake Kinneret,Israel

Lake Kinneret, Israel, is a warm (13–30°C) monomictic lake that stratifies in April and turns over in December. Between January and June each year, a heavy bloom (up to 250 g wet weight n^–2 2) of the dinoflagellate Peridinium gatunense dominates the phytoplankton biomass. In early summer, the bloom collapses, and the sinking Peridinium biomass serves as a trigger for intense sulfate-reduction activity throughout the hypolimnion and within the sediments. The availability of organic matter and sulfate was high shortly after the bloom crash and the beginning of stratification and was lowest in December before overturn. Sulfate-reduction rates at three different sites in the lake were studied. In the sediments, the rates varied seasonally and among stations from 5 to 1600 nmol SO₄ ^–2 reduced cm^–3 day^–1, with respect to the distance from the Jordan River, depth, organic content, and stratification period. During years of low lake water levels, intense sulfate reduction occurred in the hypolimnion, resulting in anoxia and high concentrations of H₂S (>400 m). In years with high water levels, early bloom, and delayed stratification, higher rates of sulfate reduction were recorded in the sediments, probably as a result of a greater fraction of the primary production (organic matter) reaching the bottom. Correspondence to: O. Hadas.     

Seasonal dynamics of picophytoplankton in Lake Kinneret, Israel

1. Picophytoplankton (picocyanobacteria and picoeukaryotes) communities in Lake Kinneret were studied from 1988 to 1992. No prochlorophytes were observed in the lake. 2. Picocyanobacteria were a prominent and ubiquitous component of the phytoplankton, being present at all depths throughout the year, with concentrations ranging from 2 ± 10–8 ± 10⁵ cells ml^?1. Low cell numbers in winter and spring were followed at the end of the annual dinoflagellate bloom by maximal abundances in summer-autumn in the epilimnion. High cell numbers (> 10⁴ cells ml^?1) were sometimes also found in the anaerobic hypolimnion. Net growth rates for picocyanobacteria ranged from 0.29 to 0.60 divisions day^?1. 3. Picoeukaryotes were a very minor constituent of the picoplankton, mostly present in winter and spring, and sometimes at the end of autumn, with concentrations ranging from 44 to 5700 cells ml^?1. Higher cell numbers tended to occur in the near surface water layers. In August-September, picoeukaryotes were found only in the hypolimnion. In December, the occurrence of picoeukaryotes in the deep water layers probably resulted from advection with cold water currents from the Jordan river. Net growth rates for picoeukaryotes ranged from 0.26 to 0.43 divisions day^?1. 4. Overall, the contribution of picophytoplankton to the phytoplankton standing crop in Lake Kinneret was limited; picocyanobacteria and picoeukaryotes accounted for no more than 7.0 and 0.1% of total algal biomass (semiannual average), respectively. 5. Picophytoplankton cell numbers in pelagic waters were usually similar to those in shallower lake stations. 6. Picocyanobacteria appear to be an autochthonous population, whereas picoeukaryotes are probably brought annually by the Jordan River and do not maintain themselves in the lake.     

The distribution of chlorophylls, carotenoids and their breakdown products in Lake Kinneret (Israel) sediments

SUMMARY. 1. The sedimentary distribution of chlorophylls, carotenoids and their breakdown products following the winter-spring bloom of the dinoflagellate Peridinium gatunense in Lake Kinneret was determined both spatially and vertically within the sediments, using high-pressure liquid chromatography (HPLC).
2. All four stations (epilimnic, oxic St. J (depth = 8m) and D (12m) and hypolimnic, anoxic St. F (21 m) A (41 m) were characterized by the presence of chlorophyll a (chl-a) chl- b , chl- c , phaeophytins a (ph'tin- a ) and b , phaeophorbide a (ph'bide- a ) and lutein, fucoxanthin, diatoxanthin and β-carotene together with spectrally similar, but HPLC-resolved, breakdown products in the surface sediments. The central deep station A (41 m), which is seasonally anoxic, trapped and preserved significantly more pigments (e.g. chl- a = 16.35 μg g⁻¹ organic matter) than the more erosive, oxic littoral stations (chl- a = 1.10 and 0.76μg g^−l OM at St. D and J, respectively).
3. Whereas it was possible to ascribe the presence of lutein and chl- b to inputs of green algae and higher plants from the catchment, and fucoxanthin and chl- c ₁, c ₂ to winter diatom blooms (e.g. Melosira granulata ), there was a remarkable absence of peridinin and dinoxanthin from sediments, despite the dominance of dinoflagellates in the phytobiomass. This is consistent with the well-known oxic lability of this epoxy carotenoid in aquatic systems, and indicates efficient breakdown of this dinoflagellate organic matter in the water column, prior to sedimentation.
4. Although pigment concentrations all decreased with depths within sediments, lutein and β-carotene were selectively preserved compared to chl-a and fucoxanthin which, at 40m depth at St. A, decreased to < 1% of their surface concentrations.     

Aphanizomenon ovalisporum (Forti) in Lake Kinneret, Israel

The filamentous cyanobacterium Aphanizomenon ovalisporum wasobserved for the first time in Lake Kinneret in August 1994and formed a prominent bloom from September through October.Aphanizomenon ovalisporum reappeared in diminished amounts inthe summer and fall of 1995. These events are the first recordof significant quantities of a potentially toxic nitrogen-fixingcyanobacterium in this lake. No definite provenance of inoculumhas been identified, although A.ovalisporum was also observedin a newly reflooded area (Lake Agmon) in the catchment. Unusuallyhigh water temperatures and low wind inputs were observed priorto and during the A.ovalisporum bloom period. These, togetherwith possibly enhanced availability of phosphorus or other growthfactors, may have contributed to the cyanobacterium growth in1994. Phosphorus limi tation, as indicated by high cellularalkaline phosphatase activity, the onset of stormy conditionsand a fall in water temperatures led to the demise of the 1994bloom. Although the A. ovalisporum bloom in 1994 had no seriousdirect impact on water quality, the continued presence of apotentially toxic cyanobacterium in Lake Kinneret, a major nationalwater supply source, is a cause for serious concern.     

Appearance and establishment of diazotrophic cyanobacteria in Lake Kinneret,Israel

1. We propose that the appearance and establishment of Nostocales (cyanobacteria) species of the genera Aphanizomenon and Cylindrospermopsis in the warm subtropical Lake Kinneret (Sea of Galilee, Israel) from 1994 was linked to changes in climate conditions and summer nitrogen (N) availability. 2. From 1994 to 2009, an increase in frequency of events of elevated water temperature (>29 °C) in summer, and to some extent a greater frequency of lower summer wind speed events, affected water turbulence and water column stratification, thus providing better physical conditions for the establishment of these populations. 3. In recent years, N‐depleted conditions in Lake Kinneret in early summer have promoted the development and domination of Nostocales that could gain an ecological advantage owing to their N₂‐fixing capability. 4. Nitrogen fixation rates coincided both with heterocyst abundance and with Nostocales biomass. The N supplied to the lake via nitrogen fixation ranged from negligible quantities when Nostocales represented only a minor component of the phytoplankton community to 123 tonnes when Cylindrospermopsis bloomed in 2005. This high N₂ fixation rate equals the average summer dissolved inorganic nitrogen load to the lake via the Jordan River.     

Seasonal photosynthetic activity of autotrophic picoplankton in Lake Kinneret, Israel

Autotrophic picoplankton populations in Lake Kinneret are composedof picocyanobacteria and picoeukaryotes. Overall, the ratesof photosynthetic carbon fixed by autotrophic picoplankton duringthis study were low (0.01–1.5 mg Cm^–3 h^–1).The highest chlorophyll photosynthetic activity of the <3µm cell-size fraction was found in spring, when picoeukaryotespredominated and in addition small nanoplankton passed throughthe filters. The maximum cell-specific photosynthetic rate ofcarbon fixation by picocyanobacteria and picoeukaryotes was2.5 and 63 fg C cell^–1 h^–1, respectively. The highestspecific carbon fixation rate of autotrophic picoplankton was11 µg C µg^–1 Chl h^–1 The proportionalcontribution of autotrophic picoplankton to total photosynthesisusually increased with depth. Picocyanobacteria collected fromthe dark, anaerobic hypolimnion were viable and capable of activephotosynthesis when incubated at water depths within the euphoticzone. Maximum rates of photosynthesis (P_max) for picocyanobacteriaranged from 5.4 to 31.4 fg C cell^–1 h^–1 with thehighest values in hypolimnetic samples exposed to irradiance.Photosynthetic efficiency (     

Decreased salinity effects in Lake Kinneret (Israel)

Kinneret is the only freshwater lake in Israel. It currently supplies about 30% of national water demands. Most pumped water is for drinking, and water quality is of major concern. During 1970–1987 temporal changes were observed in the lake ecosystem: decrease of salinity, decrease of total N (TN) and increase of total P (TP) mass contents, decline of TN/TP ratio, increase of phytoplankton biomass and increase of algal photosynthetic specific activity. It is suggested that because of decrease in salinity carbonic anhydrase activities in algal cells and nitrifying bacteria were enhanced. The increase of NO₃ flux through nitrification consequently enhanced denitrification and nitrogen losses in lake waters. These increased N losses together with P increase, as reflected by the decline of TN/TP ratio might be a slight shift from the present both P and N deficiencies to a higher level of N limitation in the Kinneret ecosystem. This This may cause changes in phytoplankton community structure possibly without changing primary production levels but deteriorate water quality.     

Phytoplankton periodicity in a subtropical lake (Lake Kinneret,Israel)

Lake kinneret is a subtropical monomictic lake characterized by a Pyrrhophyta-Chlorophyta assemblage, supplemented by Cyanophyta in some years. Concerning their abundance and seasonal occurrence, the phytoplanktonic algae belong to two groups: algae appearing in quantity at a definite annual period and algae present throughout the year. Four stages of algal succession occur in the lake. There is a marked periodicity in the phytoplankton composition with a high standing stock in winter-spring, due to the dinoflagellate water-bloom, and a low one during the summer months, related to the high stability of summer stratification. The annual succession at the species level has been an almost constant event in the lake for many years.The increase in nutrient concentrations in 1973 and 1974 increased the diversity and abundance of algae (except Peridinium) but did not lead to significant changes in algal succession. Conversely, the decrease of the zooplankton grazing pressure in 1975 and 1976 facilitated the development of algal maxima during summer-fall. They were caused by nanoplanktonic forms, and they developed without additional enrichment of nutrients. The algal abundance and diversity decreased. The years 1981 and 1982 were characterized by both an increase in phosphorus and a decrease in zooplankton. These conditions favored the concomitant abundance of many species and an increase of non-Pyrrhophyta biomass.     

Acoustic assessment of the number of pelagic fish in Lake Kinneret,Israel

The pelagic fish stock (mostly lavnun, Mirogrex terraesanctae) of Lake Kinneret was assessed using HADAS, a newly developed echo-counting analysis system. Total fish numbers (during Mar–Apr) were estimated to be 61 million in 1987, 218 million in 1988, 120 million in 1989, and 69 million in April 1990. The total fish number for March 1981 was recalculated from published data and estimated at 170 million. The decline in total numbers between 1988 and 1990 resulted primarily from a decrease in the relative abundance of the smallest size classes of fish. Since fish abundance apparently declined between 1981 and 1990, the decline in zooplankton standing stocks documented during these years cannot be attributed to higher fish predation caused by larger fish stocks. The HADAS estimates of lavnun population size are large enough to account for the known catch of lavnun.Contribution from Israel Oceanographic & Limnological Research, Ltd. This research was supported in part by the Israeli Water Commissioner, the Israeli Electric Company, and by a grant from the U.S. Agency for International Development.     

High abundance of picoplankton-ingesting ciliates during late fall in Lake Kinneret, Israel

Small, aloricate ciliates dominated the biomass of heterotrophicprotists throughout the water column at the end of the periodof stratification in Lake Kinneret, Israel The integrated biomassof cilates was 5–20 times that of heterotrophic flagellatesDuring incubation experiments, ciliate growth rates in cpilimneticwater corresponded to population doubling times of 9.6–19.4h, while flagellate populations showed no growth. Most of thealiates were small forms (10–30 µm long), includingscuticocihates, choreotnchs, Coleps spp. and Colpoda spp., andappeared to be consuming bacteria, coccoid cyanobacteria, and<5 µm eukaryotic algae. Grazing rates of cihate assemblageson picoplankton in the epilimnion, as determined by the uptakeof fluorescently labeled bacteria and cyanobactena, ranged from62 to 86 nl cell^–1 h^–1 Colpoda steini, isolatedfrom lakewater, grew on a cultured freshwater Synechococcussp with a doubling time of 4.5 h, and a gross growth efficiencyof 48% The estimated daily requirements of ciliates for growthapproximately equalled total phytoplankton production. We calculatedthat ciliates in the epilimnion were clearing 4–10% ofthe bacterioplankton and cyanobactenal standing stocks per daySince this would not be sufficient food consumption to meetdaily carbon requirements of the aliates, it is likely thatthese organisms were also grazing a significant amount of autotrophicand heterotrophic eukaryotic cells in Lake Kinneret.     

Limnological and ecophysiological aspects of Aphanizomenon ovalisporum bloom in Lake Kinneret, Israel

The first appearance of Aphanizomenon ovalisporum in Lake Kinneret in August 1994 was apparently boosted by relatively high concentrations of total dissolved phosphorus (12 g P l^-1 as compared to an average of 8 g P l^-1). The increasing Aphanizomenon biomass in a lake in which phytoplankton are generally phosphate limited in summer and autumn was accompanied by high enzymatic activity of alkaline phosphatase, reaching values of 2830 nmol MU l^-1 h^-1, suggesting a great demand for phosphorus. In addition, the nitrogen requirement of the developing population of Aphanizomenon was partly provided by nitrogen fixation, as indicated by a high percentage of heterocysts. Laboratory experiments demonstrated that filtrate from an old Peridinium gatunense culture enhanced Aphanizomenon growth. Thus, it is postulated that the degradation of the massive Peridinium bloom in spring and early summer supported the development of A.ovalisporum. The high pH and alkalinity during the bloom of Aphanizomenon indicate that A.ovalisporum is probably a HCO3^- user. After 1994, akinetes of A.ovalisporum were left in sediments and the water column, and could be a source for the next year's bloom. This possibility was demonstrated by inoculation of lake water and sediments into nitrogen-depleted BG-11 medium, resulting in the dominance of A.ovalisporum.      

The structure and dynamics of the phytoplankton assemblages in Lake Kinneret, Israel

The phytoplankton of Lake Kinneret, a warm monomictic lake,is dominated by a Pyrrhophyta-Chlorophyta assemblage. Four stagesof succession of planktonic algae occur in the lake, startingwith thermal and chemical destratification and ending with stratification. The index of diversity of the phytoplankton communities is highduring the destratification and mixed periods. The index reachesminimal values during late summer, when the ecosystem is subjectto strong physical, chemical and biological stresses. The diversityin Lake Kinneret increases with the increase in nutrients andnot with the increase in temperature. During most of the year, the nanoplanktonic forms are in greaternumbers than the netplankton species. This fact is correlatedwith the amounts of available nutrients in the lake. The annual averages of the wet autotrophic biomass in Lake Kinneretare very high in comparison with other warm lakes. The contributionof the nanoplanktonic species to the total algal biomass isvery small during the Peridinium bloom, but represents approximatelyhalf of the total algal biomass during the rest of the year. The concentration of nutrients in the water, together with theadverse competitive effect of Peridinium on other algae, areto a large extent responsible for the composition, successionand abundance of the phytoplankton assemblages in Lake Kinneret.     

Eukaryotic picoplankton, Mychonastes homosphaera (Chlorophyceae, Chlorophyta), in Lake Kinneret, Israel

Eukaryotic picoplankton was isolated from Lake Kinneret, Israel. This green picoplankton is spheroidal or ovoidal in shape. The cell wall has a network of ribs on its surface. The chloroplast is cup- or mantel-shaped without a pyrenoid. The morphology and cellular structure of this autosporic alga is identical with Mychonastes homosphaera (Skuja) Kalina et Pun?ochá?ová. The phylogenetic tree based on 18S ribo-somal RNA gene sequence data showed that M. homosphaera is placed in the Chlorophyceae and located at the base of a branch leading to the other algae possessing directly opposed orientation of the flagellar apparatus.     

Release of dissolved organic matter by photosynthesizing algae in Lake Kinneret,Israel*

The incorporation of inorganic carbon in particulate and dissolved organic matter by natural populations of photosynthesizing algae has been studied in Lake Kinneret, Israel, from August 1972 to December 1973. The release of dissolved organic material was directly related to the rates of particulate productivity. However, the percentage of extracellular release (PER) of dissolved organic compounds showed an inverse relationship with chlorophyll a concentrations. Mean PER values (for the trophogenic layer) over the experimental period were 3·7%, and were lowest during the exponential growth phase of the annual dinoflagellate bloom (mean PER = 1·9%). Highest PER were measured in deep, light-limited samples (mean = 31·7%; s.d.· 21·9%). In a dense metalimnic layer of photosynthetic sulphur bacteria (Chlorobium phaeobacterioides), PER was not as extremely high as has been reported for a similar layer in Lake Wadolek, Poland. Although the amounts of dissolved organic matter released by phytoplankton in Lake Kinneret were low, they may have significance as growth modulators and as substrates for heterotrophic organism.     

Simultaneous patterns of temperature, oxygen, zooplankton and fish distribution in Lake Kinneret, Israel

1. Temperature, oxygen concentration and zooplankton density were measured simultaneously on 25 December 1990 at twenty-six stations during an acoustic survey of the fish population of Lake Kinneret, Israel. Data from a preliminary acoustic survey were used to establish the optimal distance between transects (4km). 2. The distribution of temperature and oxygen concentration (and their gradients), and zooplankton and fish densities were compared using three-dimensional plots and correlation analysis. Both fish and zooplankton were concentrated in frontal zones where temperature and oxygen gradients were highest. 3. In the south-eastern part of the lake, where fish abundance was highest, a minimum of zooplankton density was observed, suggesting that predation by fish reduced zooplankton concentrations.     

Alternative states in the phytoplankton of Lake Kinneret, Israel (Sea of Galilee)

1. Through analyses of a 34‐year record of phytoplankton, zooplankton and physicochemical parameters from Lake Kinneret, Israel, we show that distinct and persistent phytoplankton assemblage states occurred from winter to summer. 2. The most obvious characteristic of these states was the presence or absence of a spring bloom of the dinoflagellate, Peridinium gatunense. 3. Analyses of the data within the framework of the alternative states model revealed a possible complex triggering mechanism, and system hysteresis. 4. A change in zooplankton biomass and body size coincident with changes in predation pressure associated with the collapse of the Kinneret Bleak, Acanthobrama terraesanctae, fishery appeared to be the ‘slow changing’ variable in the context of the alternative states model. Alternative phytoplankton states were only possible after this variable crossed a threshold in 1993–94, following the collapse of the fishery. 5. When alternative states were possible, some physicochemical parameters and the structure of the zooplankton assemblage appeared to control which phytoplankton state emerged in a given year. In years without a P. gatunense bloom, important physicochemical parameters in winter included low NO₃ loading, high water temperature, high water level, a deeper thermocline, low transparency, high concentrations of NO₃ and Cl in the epilimnion, and low concentration of epilimnetic total phosphorus. In addition, the cladoceran Chydorus sphaericus and adults of the copepod Mesocyclops ogunnus were observed in winter in years without a bloom. 6. Zooplankton biomass and body size of some taxa have recovered since the 1993–94 collapse of the fishery, yet incidence of both phytoplankton states in Lake Kinneret was still possible. Within the framework of the alternative states model, this suggests that the slow changing variable threshold where alternative states became possible is different from the threshold where alternative states will no longer be possible. In other words, the system is characterised by a hysteresis.